Keith Jensen

What's in it for me? Self-regard precludes altruism and spite in chimpanzees

Sensitivity to fairness may influence whether individuals choose to engage in acts that are mutually beneficial, selfish, altruistic, or spiteful. In a series of three experiments, chimpanzees (Pan troglodytes) could pull a rope to access out-of-reach food while concomitantly pulling another piece of food further away. In the first study, they could make a choice that solely benefited themselves (selfishness), or both themselves and another chimpanzee (mutualism). In the next two experiments, they could choose between providing food solely for another chimpanzee (altruism), or for neither while preventing the other chimpanzee from receiving a benefit (spite). The main result across all studies was that chimpanzees made their choices based solely on personal gain, with no regard for the outcomes of a conspecific. These results raise questions about the origins of human cooperative behaviour.

Chimpanzees are vengeful but not spiteful

People are willing to punish others at a personal cost, and this apparently antisocial tendency can stabilize cooperation. What motivates humans to punish noncooperators is likely a combination of aversion to both unfair outcomes and unfair intentions. Here we report a pair of studies in which captive chimpanzees (Pan troglodytes) did not inflict costs on conspecifics by knocking food away if the outcome alone was personally disadvantageous but did retaliate against conspecifics who actually stole the food from them. Like humans, chimpanzees retaliate against personally harmful actions, but unlike humans, they are indifferent to simply personally disadvantageous outcomes and are therefore not spiteful.

Punishment and spite, the dark side of cooperation

Causing harm to others would hardly seem to be relevant to cooperation, other than as a barrier to it. However, because selfish individuals will exploit cooperators, functional punishment is an effective mechanism for enforcing cooperation by deterring free-riding. Although functional punishment can shape the social behaviour of others by targeting non-cooperative behaviour, it can also intimidate others into doing almost anything. Second-party functional punishment is a self-serving behaviour at the disposal of dominant individuals who can coerce others into behaving cooperatively, but it need not do so. Third-party and altruistic functional punishment are less likely to be selfishly motivated and would seem more likely to maintain norms of cooperation in large groups. These forms of functional punishment may be an essential part of non-kin cooperation on a scale exhibited only by humans. While punitive sentiments might be the psychological force behind punitive behaviours, spiteful motives might also play an important role. Furthermore, functionally spiteful acts might not be maladaptive; reckoning gains relative to others rather than in absolute terms can lead to hyper-competitiveness, which might also be an important part of human cooperation, rather than just an ugly by-product.

The emergence of human prosociality: aligning with others through feelings, concerns, and norms

The fact that humans cooperate with nonkin is something we take for granted, but this is an anomaly in the animal kingdom. Our species’ ability to behave prosocially may be based on human-unique psychological mechanisms. We argue here that these mechanisms include the ability to care about the welfare of others (other-regarding concerns), to “feel into” others (empathy), and to understand, adhere to, and enforce social norms (normativity). We consider how these motivational, emotional, and normative substrates of prosociality develop in childhood and emerged in our evolutionary history. Moreover, we suggest that these three mechanisms all serve the critical function of aligning individuals with others: Empathy and other-regarding concerns align individuals with one another, and norms align individuals with their group. Such alignment allows us to engage in the kind of large-scale cooperation seen uniquely in humans.

Restorative Justice in Children

An important, and perhaps uniquely human, mechanism for maintaining cooperation against free riders is third-party punishment. Our closest living relatives, chimpanzees, will not punish third parties even though they will do so when personally affected. Until recently, little attention has been paid to how punishment and a sense of justice develop in children. Children respond to norm violations. They are more likely to share with a puppet that helped another individual as opposed to one who behaved harmfully, and they show a preference for seeing a harmful doll rather than a victim punished. By 6 years of age, children will pay a cost to punish fictional and real peers, and the threat of punishment will lead preschoolers to behave more generously. However, little is known about what motivates a sense of justice in children. We gave 3- and 5-year-old children--the youngest ages yet tested--the opportunity to remove items and prevent a puppet from gaining a reward for second- and third-party violations (experiment 1), and we gave 3-year-olds the opportunity to restore items (experiment 2). Children were as likely to engage in third-party interventions as they were when personally affected, yet they did not discriminate among the different sources of harm for the victim. When given a range of options, 3-year-olds chose restoration over removal. It appears that a sense of justice centered on harm caused to victims emerges early in childhood and highlights the value of third-party interventions for human cooperation.

The nature of prosociality in chimpanzees

An important debate centres around the nature of prosociality in nonhuman primates. Chimpanzees help other individuals in some experimental settings, yet they do not readily share food. One solution to this paradox is that they are motivated to help others provided there are no competing interests. However, benefits to recipients could arise as by-products of testing. Here we report two studies that separate by-product from intended helping in chimpanzees using a GO/NO-GO paradigm. Actors in one group could help a recipient by releasing a food box, but the same action for another group prevented a recipient from being able to get food. We find no evidence for helping—chimpanzees engaged in the test regardless of the effects on their partners. Illusory prosocial behaviour could arise as a by-product of task design.

Chimpanzee responders still behave like rational maximizers

The ultimatum game (1) is a powerful and widely used test of bargaining behavior that has only recently been applied to nonhuman animals (2⇓–4). The key feature of this game is the power the responder has; the threat of rejections—the ultimatum—typically induces proposers to be more generous than they would be otherwise. Proctor et al. (4) suggested that chimpanzees exhibit sensitivity to fairness in a more “intuitive” ultimatum game based on token exchange, contrary to refs. 2 and 3. This contradiction, however, is more apparent than real. Responders in Proctor et al. (4) accepted 100% of all offers, even more than chimpanzees and bonobos in …

Large-scale cooperation

A deeper understanding of the evolution of cooperation will come from investigations of what animals know about working together. A study with Asian elephants now adds to the literature on the subject.

Preschoolers are sensitive to free riding in a public goods game

Despite the benefits of cooperation, selfish individuals often produce outcomes where everyone is worse off. This “tragedy of the commons” has been demonstrated experimentally in adults with the public goods game. Contributions to a public good decline over time due to free-riders who keep their endowments. Little is known about how children behave when confronted with this social dilemma. Forty-eight preschoolers were tested using a novel non-verbal procedure and simplified choices more appropriate to their age than standard economic approaches. The rate of cooperation was initially very low and rose in the second round for the girls only. Children were affected by their previous outcome, as they free rode more after experiencing a lower outcome compared to the other group members.

Correspondence: Reply to ‘Chimpanzee helping is real, not a byproduct’

J.C. was supported in part by the European Research Council under the European Union’s Seventh Framework Programme (FP7/2007-2013)/ERC grant agreement no. 609819, project SOMICS.